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Population Comparison of Right Whale Body Condition Reveals Poor State of the North Atlantic Right Whale

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Population Comparison of Right Whale Body Condition Reveals Poor State of the North Atlantic Right Whale Vol. 640: 1–16, 2020 MARINE ECOLOGY PROGRESS SERIES Published April 23 https://doi.org/10.3354/meps13299 Mar Ecol Prog Ser OPENPEN ACCESSCCESS FEATURE ARTICLE Population comparison of right whale body condition reveals poor state of the North Atlantic right whale Fredrik Christiansen1,2,3,*, Stephen M. Dawson, John W. Durban, Holly Fearnbach, Carolyn A. Miller, Lars Bejder, Marcela Uhart, Mariano Sironi, Peter Corkeron, William Rayment, Eva Leunissen, Eashani Haria, Rhianne Ward, Hunter A. Warick, Iain Kerr, Morgan S. Lynn, Heather M. Pettis, Michael J. Moore 1Aarhus Institute of Advanced Studies, Høegh-Guldbergs Gade 6B, 8000 Aarhus C, Denmark 2Zoophysiology, Department of Biology, Aarhus University, C.F. Møllers Alle 3, 8000 Aarhus C, Denmark 3Centre for Sustainable Aquatic Ecosystems, Harry Butler Institute, Murdoch University, Murdoch, 6150 Western Australia, Australia The complete list of author addresses is in the Appendix ABSTRACT: The North Atlantic right whale Eubala - ena glacialis (NARW), currently numbering <410 individuals, is on a trajectory to extinction. Al though direct mortality from ship strikes and fishing gear entanglements remain the major threats to the popu- lation, reproductive failure, resulting from poor body condition and sublethal chronic entanglement stress, is believed to play a crucial role in the population decline. Using photo grammetry from unmanned aer- ial vehicles, we conducted the largest population assessment of right whale body condition to date, to determine if the condition of NARWs was poorer than 3 seemingly healthy (i.e. growing) populations of southern right whales E. australis (SRWs) in Argen - Three healthy southern right whales (left three photographs) tina, Australia and New Zealand. We found that next to a North Atlantic right whale (right) in visibly poorer body condition NARW juveniles, adults and lactating females all had lower body condition scores compared to the SRW Photos: Fredrik Christiansen (left & center-left), Stephen M. Dawson (center-right), John W. Durban/Holly Fearnbach (right) populations. While some of the difference could be the result of genetic isolation and adaptations to local environmental conditions, the magnitude suggests KEY WORDS: Baleen whale · Bioenergetics · that NARWs are in poor condition, which could be Eubalaena · Morphometrics · Photogrammetry · suppressing their growth, survival, age of sexual Unmanned aerial vehicles maturation and calving rates. NARW calves were found to be in good condition. Their body length, however, was strongly determined by the body con- dition of their mothers, suggesting that the poor condition of lactating NARW females may cause a 1. INTRODUCTION reduction in calf growth rates. This could potentially lead to a reduction in calf survival or an in crease in Right whale populations around the world were se- female calving intervals. Hence, the poor body con- verely depleted (some reduced to less than 5% of the dition of individuals within the NARW population is original population size) by commercial whaling op- of major concern for its future viability. erations from the 11th to the 20th century in the North Atlantic (Aguilar 1986, Reeves et al. 1999) and during © The authors 2020. Open Access under Creative Commons by *Corresponding author: [email protected] Attribution Licence. Use, distribution and reproduction are un - restricted. Authors and original publication must be credited. Publisher: Inter-Research · www.int-res.com 2 Mar Ecol Prog Ser 640: 1–16, 2020 the 19th to 20th century in the Southern hemisphere (Christiansen et al. 2016a). Like most baleen whales, (Dawbin 1986, Jackson et al. 2016), as well as by ille- right whales make annual migrations between high- gal Soviet whaling in the Southern hemisphere in the latitude feeding grounds in summer and low-latitude 1950s to early 1970s (Yablokov 1994, Tormosov et al. breeding grounds in winter (Bannister et al. 1999). 1998). Since the cessation of commercial whaling, the Females become sexually mature at around 9 yr old southern right whale Eubalaena australis (SRW) has and give birth to a single calf at a time (Kraus et al. been recovering at a relatively rapid pace throughout 2001, Cooke et al. 2003, Burnell 2008). They are ‘cap- most of its range, and currently numbers in the tens ital’ breeders, fasting during the winter breeding of thousands globally (IWC 2013). The population season, and thus have a finite amount of energy to growth rate during this time has been as high as 5.55% invest in late pregnancy and lactation (Lockyer 1987, for Australia (Bannister 2016), 5−7% for New Zealand Stephens et al. 2009). Christiansen et al. (2018) (Carroll et al. 2013) and 6.5% for Argentina (Cooke et showed in SRWs that maternal size (body length and al. 2015), although the growth rate of the latter has condition) has a direct effect on the amount of energy slowed down substantially (Crespo et al. 2019). that lactating females invest in their calves, which in In contrast, the recovery of the North Atlantic right turn dictates calf growth rates. When conditions are whale E. glacialis (NARW) has been considerably favourable, females generally have a 3 yr reproduc- slower, with a mean annual growth rate of 2.8% be- tive cycle consisting of 1 yr of gestation, 1 yr of lacta- tween 1990 and 2010 (Pace et al. 2017). More recent tion and 1 yr of resting (to recover energy stores) abundance estimates, between 2010 and 2015, indicate (Best 1994). The mean calving interval for SRWs is that the population has been declining at a rate of just close to this 3 yr minimum, at 3.33 yr in Australia under 1% per year (Pace et al. 2017). The rate of (Burnell 2001), 3.31 yr in New Zealand (Davidson et decline has been higher for females, which dropped at al. 2017) and between 2.96 and 3.24 yr in Argentina approximately 7% between 2010 and 2015, compared (Marón et al. 2015). In contrast, since 2015, the mean to about 4% for males over the same period (Pace et al. calving interval for NARW females is >7 yr (Pettis et 2017). The situation for NARWs was further worsened al. 2020), suggesting that they need several years by an unusual mortality event between 1 November longer to recover from a reproductive event. Apart 2016 and 31 December 2017, when at least 17 juvenile from body condition having a direct effect on female and adult right whales died as a result of entangle- reproductive success, it can also influence juvenile ments and vessel strikes (NARWC 2018). In December growth rates (Douhard et al. 2017) and the age of 2015, prior to the 2017 mortalities, the species’ abun- sexual maturation (Sigurjónsson et al. 1990), which dance was estimated at 451 individuals, of which 186 could negatively influence population growth. were females. The best estimate as of the end of 2017 The aim of this study was to assess the body condi- was 411 animals (Pettis et al. 2018). While fishing gear tion of the NARW. Although the population’s body entanglements and ship strikes are the largest direct condition (based on visual assessment) has declined anthropogenic threats to the NARW population during the last 3 decades (Rolland et al. 2016), a com- (Moore et al. 2004, Knowlton et al. 2012, van der Hoop parison to healthy (growing) right whale populations et al. 2013, Kraus et al. 2016), reduced reproductive is needed to assess its current status. Unfortunately, rate resulting from nutritional stress (i.e. poor body no historical data on NARW body condition exist to condition) has been hypothesised as a factor further allow such a comparison. Instead, the best opportu- contributing to the population decline (Kraus et al. nity to assess the relative body condition of NARW 2001, Reeves et al. 2001, Schick et al. 2013, Rolland et comes from a comparison with their closest living rel- al. 2016). The sublethal impacts of entanglement have ative, the SRW. We therefore compared the body con- also been modelled to significantly impact reproduc- dition of NARWs with 3 seemingly healthy (i.e. grow- tive success (van der Hoop et al. 2017). ing) populations of SRW in Australia, New Zealand The effect of body condition on reproduction is well and Argentina. Although we were comparing 2 dif- documented in both terrestrial (Albon et al. 1983, ferent species of right whales (Rosenbaum et al. 2000, Loudon et al. 1983, Skogland 1984, Atkinson & Gaines et al. 2005), which might differ in their body Ramsay 1995, Festa-Bianchet 1998) and marine condition due to genetic differences, our rationale mammals (Arnbom et al. 1997, Boltnev & York 2001, was that body condition, similar to most traits closely Bowen et al. 2001, Wheatley et al. 2006). In baleen associated with fitness, shows low genetic variance whales, female body condition influences fecundity relative to environmental variance (Mousseau & Roff (Lockyer 2007, Williams et al. 2013), foetal growth 1987, Kruuk et al. 2000). We also show that NARWs (Christiansen et al. 2014) and calf body condition and SRWs are very similar in body shape, size and life Christiansen et al.: Body condition of right whale populations 3 history characteristics, which should facilitate com- North Atlantic, Australia, New Zealand and Argen - parison. Based on the lower population growth rate tina (Fig. 1). Photographs of NARW lactating females and longer calving interval of the NARW, our main and calves were collected on their calving grounds in hypothesis is that NARWs are in poorer body condi- Florida, USA, between 12 January and 22 February tion compared to SRWs. To help infer the potential ef- 2016 and 2017, while juvenile and adult NARWs fects of reduced body condition on different life his- were photographed on their feeding grounds in Cape tory parameters, we split our analysis into different Cod Bay, USA, between 21 March and 27 April 2016 reproductive classes (calves, juveniles, adults and and 2017 (Fig.
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